Phylogeny of African Myotis Bats (Chiroptera, Vespertilionidae) Inferred from Cytochrome B Sequences
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Acta Chiropterologica, 6(2): 177–192, 2004 PL ISSN 1508-1109 © Museum and Institute of Zoology PAS Phylogeny of African Myotis bats (Chiroptera, Vespertilionidae) inferred from cytochrome b sequences BENOÎT STADELMANN1, 2, DAVID S. JACOBS3, CORRIE SCHOEMAN3, and MANUEL RUEDI1, 4 1Natural History Museum, P.O. Box 6434, 1211 Geneva 6, Switzerland 2Department of Zoology and Animal Biology, Molecular Systematics Group, University of Geneva, 30 quai Ernest-Ansermet, CH-1211 Geneva 4, Switzerland 3Department of Zoology, University of Cape Town, Private Bag, Rondebosch 7701, South Africa 4Corresponding author: [email protected] The genus Myotis is comprised of about 100 species that are unequally distributed between the Northern (81% of the species) and the Southern hemisphere (19% of the species). Only eight species of Myotis occur in the Ethiopian region, but this is the only biogeographic region with representatives of all four classical subgenera, suggesting a diverse assemblage of morphotypes. We used sequences of a mitochondrial DNA gene (cyt b) to investigate the evolution and the phylogenetic position of seven of the eight Ethiopian species, and compared them to a broad sampling of Myotis from the World and of other vespertilionids. Phylogenetic reconstruction was based on 91 complete sequences representing 79 species of bats. The two endemic southern African species of the subgenus Cistugo were not placed within the genus Myotis, but were basal to the vespertilionid radiation, as suggested by earlier work based on karyology. The remaining Ethiopian species formed a strong mono- phyletic clade within Myotis, further stressing the importance of biogeography as a good predictor of phyloge- netic relationships. This Ethiopian clade includes one Western Palaearctic and one Oriental species, both of which probably secondarily colonized these areas from the Ethiopian region. Molecular dating based on Bayesian inferences suggest that these faunal exchanges occurred at the end of the Miocene, while the split of the Ethiopian clade from the other Old World Myotis dates back to the middle Miocene, quite early in the Myotis radiation. Thus, the relative paucity of species in sub-Saharan Africa cannot be attributed to a late entry into this continent. Instead, these molecular results suggest that other evolutionary processes are responsible for the poor species diversity of Myotis found in Africa today. Key words: Cistugo, African Myotis, Vespertilionidae, cytochrome b, molecular dating INTRODUCTION (55 species), followed by the Nearctic re- gion (24 species), while only 14 species are With about 100 currently recognized found in the Neotropics, eight in the species (Koopman, 1994; Simmons, In Ethiopian, and three in the Australian region press), bats of the genus Myotis represent (Koopman, 1994; Horá…ek et al., 2000). one of the few mammalian groups with a This pattern is particularly surprising since natural distribution covering most of the Myotis bats have adapted to diverse habi- world. The peak diversity of Myotis species tats, from deserts to cloud forests, with a is, however, reached in the northern conti- wide range of foraging habits (from insecti- nents. Eurasia supports the richest diversity vorous aerial hawkers to fish-eating). They Acta Chiropterologica, 6(2): 193–217, 2004 PL ISSN 1508-1109 © Museum and Institute of Zoology PAS Systematic status of African populations of Pipistrellus pipistrellus complex (Chiroptera: Vespertilionidae), with a description of a new species from Cyrenaica, Libya PETR BENDA1, PAVEL HULVA2, and JIÌÍ GAISLER3 1Department of Zoology, National Museum (Natural History), Václavské nám. 68, 115 79 Praha 1, Czech Republic; E-mail: [email protected] 2Department of Zoology, Charles University, Vini…ná 7, 128 44 Praha 2, Czech Republic 3Department of Zoology and Ecology, Masaryk University, KotláÍská 2, 611 37 Brno, Czech Republic The distribution of pipistrelles of the Pipistrellus pipistrellus complex (= P. pipistrellus s.l.) reaches only marginally the African continent. These bats are known only from a narrow belt of the Mediterranean zone in Maghreb and from NE Libya. We analysed museum specimens of African populations of P. pipistrellus s.l. using both morphologic and genetic techniques and compared them with Eurasian specimens of the complex. The African representatives of P. pipistrellus complex include two morphologically, genetically and geographically distinct populations. One of them inhabits the Mediterranean part of Cyrenaica, Libya. Belonging to the P. pygmaeus genetic lineage, these bats are represented by larger and more rusty coloured individuals with large massive rostrum and canines. In morphologic traits, this population differs significantly from all Western Palaearctic populations of the P. pipistrellus complex. These bats differ by about 6–7% in genetic distance from P. pygmaeus s. str. Within the P. pygmaeus lineage Libyan bats seem to be unique in their echolocation calls: the maximum energy of terminal frequencies was recorded at about 45 kHz. We consider the Libyan pipistrelles to represent a separate species, Pipistrellus hanaki sp. nov. Another distinct African pipistrelle population inhabits the Mediterranean parts of NW African countries, Morocco, Algeria and Tunisia. Individuals from the latter population are small and somewhat darker members of the P. pipistrellus genetic lineage, with relatively short and narrow mesial part of rostrum. Although both morphological and genetic differences between this population and Eurasian P. pipistrellus s. str. were found (genetic distance about 3–5%), they are probably not sufficient for the separation of this form at the specific level. However, the differences from European samples show rather not a cline character and therefore potential subspecific level of NW African P. pipistrellus has to be taken into consideration. Key words: Pipistrellus pipistrellus, Pipistrellus pygmaeus, Africa, morphology, morphometry, cytochrome b, mitochondrial DNA, taxonomy INTRODUCTION East up to Central Asia (Corbet, 1978; Koop- man, 1994; Taake and Vierhaus, 2004). Dis- Pipistrelle bats of the Pipistrellus pipi- tribution range reaches only the northern- strellus complex (or P. pipistrellus sensu la- most margin of the African continent, where to) are widely distributed in the western part these bats are known solely from the narrow of Eurasia, from Iberia, British Islands and belt of the Mediterranean zone in Maghreb southern Scandinavia through the Middle and from NE Libya (Fig. 1). Acta Chiropterologica, 6(2): 219–226, 2004 PL ISSN 1508-1109 © Museum and Institute of Zoology PAS A new species of Kerivoula (Chiroptera: Vespertilionidae) from Myanmar (Burma) PAUL J. J. BATES1, MATTHEW J. STRUEBIG2, STEPHEN J. ROSSITER2, TIGGA KINGSTON3, SAI SEIN LIN OO4, and KHIN MYA MYA 4 1Harrison Institute, Centre for Systematics and Biodiversity Research, Bowerwood House, St. Botolph’s Road, Sevenoaks, Kent, TN13 3AQ, United Kingdom; E-mail: [email protected] 2School of Biological Sciences, Queen Mary, University of London, London, E1 4NS, United Kingdom 3Department of Geography, Boston University, 675 Commonwealth Ave, Boston, MA 02215, USA 4Department of Zoology, University of Mandalay, Mandalay, Myanmar A new species of Kerivoula is described. Currently, it is known only from Namdee Forest in southern Kachin State, Myanmar. Externally, superficially similar to Kerivoula papillosa and with a dentition comparable to that of Kerivoula lenis, it is distinguished by its flattened skull. It was collected in evergreen forest in an area that also includes some mixed deciduous forest, shifting cultivation, and bamboo groves. Key words: Kerivoula sp. nov., Kachin State, Myanmar (Burma) INTRODUCTION pers. comm.). This team, with its experience of working in forest habitats elsewhere in In November 1999, the Harrison Insti- South-east Asia, had a particular interest in tute and the University of Yangon conduct- the forest bats of northern Myanmar. ed the first international field survey for Results of their preliminary studies in bats in Myanmar for fifty-five years. Four southern Kachin showed a particularly rich years later, following the award of a Darwin bat fauna, with fourteen species collected Initiative grant, an international bat work- from two survey sites. At least one of these shop in the University of Yangon, numerous represents a previously unknown species of additional field surveys and 10 internation- Kerivoula, which is here described and al expeditions to 12 of the 14 states and di- compared to other taxa within the subfami- visions of Myanmar, the programme of bat ly Kerivoulinae, Asiatic representatives of studies continues to develop. Results have which were recently reviewed by Vanitha- been, and are being, published in a series of rani et al. (2003). scientific papers and reports (Bates et al., 2000, 2001, 2004; Pearch et al., 2003). MATERIALS AND METHODS In March 2003, a second team of UK scientists joined forces with students and Field Survey staff of the University of Mandalay to con- Namdee Forest was one of twenty-five localities duct additional researches (M. J. Struebig, in Kachin and Shan States and Mandalay and Sagaing Acta Chiropterologica, 6(2): 227–248, 2004 PL ISSN 1508-1109 © Museum and Institute of Zoology PAS A new species of Chaerephon (Molossidae) from Madagascar with notes on other members of the family STEVEN M. GOODMAN1 and SCOTT G. C ARDIFF2 1Field Museum of Natural History, 1400 South Lake Shore Drive,